Avian genitalia, particularly in waterfowl, are extremely diverse. Penis morphology varies among species, and penis length and elaboration are associated with the frequency of forced extra-pair copulations, yet the developmental mechanisms responsible for this variation are unknown. In addition, females have a small phallic structure that is homologous to the male phallus, but little is known about when or how sexual differentiation takes place. To determine whether species-specific genital morphologies and sexual differentiation occur during duck embryonic development, we characterized development from the onset of genital tubercle initiation through stages of sexual differentiation in 3 species. Pekin and Laysan ducks have long, thick penises, whereas those of Mandarin ducks are shorter and thinner. Development of the genital tubercle is similar throughout the pre-hatching period across the 3 species, suggesting that differences in penis morphology arise post-hatching. We observed that male and female phallus development is similar at early stages, but the female phallus later regresses. Then, we compared male and female genital development between ducks and chickens, which develop a non-intromittent penis. We found that external genital development in male and female chickens resembles that of female ducks, which raises the possibility that male phallus development became feminized during galliform evolution.

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